ARTICLE 75

et is in et al.

(Stadler , Ludmila 5 (Réblová

et al. et 2016). A 11 Key words: Basidiomycetes pleomorphic fungi taxonomy protected names rejected names unit nomenclature new taxa Agaricomycotina 2017), Pezizomycetes al. et 2014) and overlooked and 2014) et al. over , are over et al. 2017, 2018). However, Sordariomycetes , Roland Kirschner Code at the 2017 International 9(1): 75–90 (2018) 75–90 9(1): · FUNGUS IMA 4 2014), al. et 2014, Visagie al. et et al. 2013), Xylariaceae et (Baral (Baral Erysiphales (Braun 2013, Johnston Dothideomycetes (Rossman are evaluated , and Amy Y. Rossman Y. Amy , and Ascomycota (Rossman 10 (Samson Helicobasidium (de Beer , and Pucciniomycotina including Ustilaginomycotina , Reinhard Berndt , Reinhard 3 Eurotiales Roestelia 2016), yeasts (Daniel yeasts 2016), et al , 320 Taoyuan City, Taiwan City, Taoyuan , 320 , Marco Thines , Marco Ustilaginomycotina over Milesia, Phragmidium 9 over Uredo. In addition, eight new over , and none in and Ophiostomatales Leotiomycetes 2014), Orbiliomycetes al. et (Healy generic names in the companion paper to this one on the et al. 2013) and remaining al. 2016) as well as 2015b), progress. Generic names that follow the principle of priority whether they are sexually or asexually typified do not need to be confirmed by the Nomenclature Committee for Fungi (NCF) and General Committee of the International Botanical approve required which 57.2 Art. former The (IBC). Congress for the use of asexually typified names over sexually typified ones was removed from the Botanical Congress (Turland ermany ersity, 625 00 Brno, Czech Republic ersity,

Dominik Begerow

, and Uromyces 1 combs. nov. and , Markus Scholler 8 Pucciniomycotina Pucciniomycotina in Fungi Gymnosporangium , Ustilaginomycotina were determined to be congeneric and 2017), al. et over Peridermium, Milesina Rhodomyces, Sordariomycetes et al. 2012), it has Naiadella and Fungi. A number of , Mehrdad Abbasi , Mehrdad 2 over 2015a), Hypocreales al. et ) and recommendations for use ) and recommendations (Basidiomycota and eight sets in and Prospodium crusculum 2014, Kepler 2014, al. et Blastoderma , Mahajabeen Padamsee 7 Classicula International Code of Nomenclature Blastospora juruensis, B. subneurophyla, Cronartium bethelii, C. kurilense, C. sahoanum, (Rossman Tuberculina, Melampsorella Tuberculina, , Lisa A. Castlebury , Lisa

1 Pucciniomycotina 2016), Microascales al. (Zhang et and Yoshitaka Ono Yoshitaka

, 6 2013, Quandt 2013, al. et With the change to one scientific name for pleomorphic fungi, generic names typified by sexual and

Diaporthales Sporobolomyces over must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). You Senckenberg Gesellschaft für Naturforschung, Frankfurt (Main), Germany Senckenberg Gesellschaft für Naturforschung, Department of Botany & Plant Pathology, Oregon State University, Corvallis, OR 97333, USA Oregon State University, Department of Botany & Plant Pathology, Systematics Team, Manaaki Whenua Landcare Research, Auckland 1072, Zealand New Landcare Research, Manaaki Whenua Team, Systematics 13, D-76133 Karlsruhe, Germany Erbprinzenstr. Staatliches Museum f. Naturkunde Karlsruhe, ETH Zürich, Plant Ecological Genetics, Universitätstrasse 16, 8092 Zürich, Switzerland Genetics, Universitätstrasse 16, 8092 ETH Zürich, Plant Ecological University Sciences and Engineering, National Central Department of Biomedical Univ Faculty of Science, Masaryk Czech Collection of Microoorganisms, Mito, Ibaraki 310-8512, Japan University, Faculty of Education, Ibaraki Mycology & Nematology Genetic Diversity and Biology Laboratory, USDA-ARS, Beltsville, MD 20705, USA USDA-ARS, Beltsville, Genetic Diversity and Biology Laboratory, Mycology & Nematology G Geobotanik, ND 03/174, D-44801 Bochum, Ruhr-Universität Bochum, Purdue University, Department of Botany and Plant Pathology, West Lafayette, IN 47901, West USA; corresponding author e-mail: maime@purdue. Department of Botany and Plant Pathology, Purdue University, Submitted: 16 March 2018; Accepted: 26 March 2018; Published: 26 April 2018. Accepted: 26 March 2018; Published: 26 Article info: Submitted: 16 March 2018; combinations are made: C. yamabense, Milesina polypodii, edu Abstract: asexual morphs have been evaluated to recommend which name to use when two names represent the same genus same the represent names two when use to name which recommend to evaluated been have morphs asexual and thus compete for generic use. names In in this paper, based on their type species to determine which names are synonyms. sets Twenty-one of sexually and asexually typified names in Aregma, compete for use. Recommendations are made as to which generic name to use. In priority most is cases followed. the However, eight principle generic of names in the recommended for protection: Marvanová © 2018 International Mycological Association © 2018 International Mycological following conditions: distribute and transmit the work, under the are free to share - to copy, You Attribution: 11 8 9 10 Ustilaginomycotina Ustilaginomycotina 5 6 7 2 3 4 1 M. Catherine Aime M. Catherine Competing sexual and asexual generic names in generic names and asexual sexual Competing Non-commercial: No derivative works: Any of the above conditions can be waived if you get For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. transform, or build upon this work. may not alter, You may not use this work for commercial purposes. You Nothing in this license impairs or restricts the author’s moral rights. permission from the copyright holder. Thanatophytum accordance with the for algae,fungi, and plants (ICN, McNeill been necessary to determine which generic name be should applied when two or more generic by different names morphs of are the same typified genus. Recommendations about which generic name to use have been made primarily by specialized Subcommissions or Working Groups of the International of Commission Fungi for (ICTF), the Taxonomy each representing a different group of papers recommending generic names for use published have including been the major groups of such as With the change to one scientific name for all INTRODUCTION doi:10.5598/imafungus.2018.09.01.06 VOLUME 9 · NO. 1 (Rossman Magnaporthales Aime et al.

those generic names that do not have priority must be that cannot be correctly placed in a genus, the problem with approved by the NCF. the generic name could be noted by including quotes around Generic names of Pucciniomycotina and Ustilaginomy- the name or by indicating the temporary taxonomic placement cotina (Basidiomycota), representing sexually and asexually with “pro tem.” as has been suggested for anamorphically typified genera that compete for use have been evaluated typified species of Ustilaginomycotina (Wang et al. 2015b).

ARTICLE by the Heterobasidiomycetes Working Group and the Efforts will be made in the future to determine the correct International Subcommission on Rust Taxonomy of the ICTF. generic placement for the names currently in use in these Members of these groups have been consulted and agree asexually typified genera. With the increasingly available use with the recommendations made herein about which generic of molecular data for determining phylogenetic relationships, name to use1. The comprehensive list of sexual-asexual the need for these asexually typified names of rust fungi generic names by Wijayawardene et al. (2012) was used as the should be eliminated or, as has been done for years, species initial starting point for discovering competing basidiomycete known only from an asexual state can be placed in the genus generic names. Citations for generic names and type species most likely to be considered correct. are based on Index Fungorum. Each set of generic names Generic names used for asexual rust fungi can, however, was evaluated based on the phylogenetic placement of their be used in a non-latinized sense to describe a morphology, type species using the literature to determine whether the as illustrated in Cummins & Hiratsuka (2003). As in other names are congeneric. Then a recommendation for use is groups of fungi, for example, a species may be described as made considering such factors as priority, number of species having ascomata that are nectria-like, meaning they produce and thus number of required name changes, use outside of light- to bright-coloured, fleshy perithecia, or fusarium-like the ICN, frequency of citations in the literature and frequency meaning that they produce fusiform, multiseptate, hyaline of use based primarily on number of citations in Google conidia. Thus, one can refer to an asexual morph of a rust Scholar (GS). fungus as being aecidium-like, caeoma-like, peridermium- Here we advocate abandoning the use of many asexually like, or uredo-like. typified generic names in Pucciniales such as Aecidium, Twenty-one sets of sexual and asexual generic names Caeoma, Milesia, Peridermium, Roestelia, and Uredo for in Pucciniomycotina and eight sets of generic names in their corresponding sexually typified generic names. This is Ustilaginomycotina were determined to compete for use problematic as many species have been described using and evaluated for recommended use. Most of the generic these names. Often it is difficult to determine in which genus names recommended for use have priority. None of the a rust fungus belongs because of the lack of the telial stage. names of Ustilaginomycotina but eight generic names of However, these formal asexually typified generic names are Pucciniomycotina do not have priority and are recommended based on type species that are synonyms of species in other for protection over older names, specifically: Classicula over genera. In the case of Aecidium and Caeoma these generic Naiadella, Gymnosporangium over Roestelia, Helicobasidium names are synonyms of an earlier generic name, Puccinia. over Thanatophytum and Tuberculina, Melampsorella For Peridermium the type species is a synonym of a species over Peridermium, Milesina over Milesia, Phragmidium in Melamsporella, a later generic name that is recommended over Aregma, Sporobolomyces over Rhodomyces and for protection over Peridermium; most species described in Blastoderma, and Uromyces over Uredo. Details about Peridermium, however, belong in Cronartium. Similarly, the each set of generic names and the basis for each decision type species of the generic name Roestelia is a synonym are presented below with an S indicating a sexually typified of the type species in Gymnosporangium, an important and name and an A indicating an asexually typified name. The well-defined genus that is recommended for protection. citation for each generic name, type species, accepted name Finally, the generic name Uredo is a synonym of the later of type species, and action required are listed in Table 1 for name Uromyces, which is recommended for use for two Pucciniomycotina and Table 2 for Ustilaginomycotina. reasons. Firstly, species of Uredo belong to many different rust genera, and secondly Uromyces is a commonly used generic name. See under those generic names for more COMPETING SEXUAL AND ASEXUAL detailed discussion. GENERIC NAMES OF PUCCINIOMYCOTINA, Although the abandoned Pucciniales asexual morph names should no longer be used to describe new species, BASIDIOMYCOTA AND RECOMMENDATIONS the species already described in those names are not FOR USE invalidated and can still be used. As for all scientific names Use Basidiopycnis Oberw. et al. 2006 (S) rather than Basidiopycnides J. Reid et al. 2008 (A) 1Heterobasidiomycetes Working Group: Dominik Bergerow, Convenor; Cathie Aime, Stephan Helfer, David McLaughlin, Hai The monotypic generic name Basidiopycnis is typified by Nguyen, Maj Padamsee, Angela Schäfer, Qi-Ming Wang, and Andrey the sexual species B. hyalina, described from bark beetle Yurkov. galleries (Oberwinkler et al. 2006). Another monotypic International Subcomission on Rust Taxonomy: Cathie Aime, Chair; generic name Basidiopycnides typified by B. albertensis Mehrdad Abbasi, Gaddam Bagyanarayana, Reinhard Berndt, was described for an asexual morph also from bark beetle Cvetomir Denchev, José Dianese, Richard Hamelin, Stephan Helfer, galleries (Hausner et al. 2008). Kirschner & Oberwinkler Drew Minnis, Yoshitaka Ono, and Markus Scholler. (2009) confirmed the relationship between these types

76 IMA FUNGUS Asexual-sexual generic names in Pucciniomycotina and Ustilaginomycotina

Table 1. Recommended generic names of Pucciniomycotina among sexual-asexual names that compete for use. The recommended name ARTICLE is in bold; see text for the rationale behind these recommendations. For each generic name this list provides the author, its date and place of publication, type species of the genus, its basionym, their dates of publication and the currently accepted name, if different. The action required is indicated in the last column such as approval by the Nomenclature Committee for Fungi (NCF) for those generic names that do not have priority and thus need protection.

Recommended generic name Synonymous alternate morph generic name(s) Action required Basidiopycnis Oberw. et al. in Mycologia 98: 639. Basidiopycnides J. Reid et al. in Mycotaxon 103: 285. None. 2006. 2008. Type: B. hyalina Oberw. et al. 2006. Type: B. albertensis J. Reid et al. 2008, now regarded as Basidiopycnis hyalina Oberw. et al. 2006. Blastospora Dietel in Ann. Mycol. 6: 222. 1908. Pelastoma M. Salazar et al. in Mycologia 104: 116. None. 2012. Type: B. smilacis Dietel 1908. Type: P. juruensis (Henn.) M. Salazar et al. 2012, basionym: Uredo juruensis Henn. 1904, now regarded as Blastospora juruensis (Henn.) Aime & Rossman 2018. Camptobasidium Marvanová & Suberkr. in Mycologia Crucella Marvanová & Suberkr., Mycologia 82: 212, None. 82: 209. 1990. 1990. Type: C. hydrophilum Marvanová & Suberkr. 1990. Type: C. subtilis Marvanová & Suberkr. 1990, now regarded as Camptobasidium hydrophilum Marvanová & Suberkr. 1990. Classicula R. Bauer et al. in Mycologia 95: 757. 2003. Naiadella Marvanová & Bandoni in Mycologia 79: Protection needed by NCF. 579. 1987. Type: C. fluitans et al. 2003. Type: N. fluitans Marvanová & Bandoni 1987, now regarded as Classicula fluitans R. Bauer et al. 2003. Cronartium Fr., Observ. Mycol. 1: 220. 1815. Endocronartium Y. Hirats. in Canad. J. Bot. 47: 1493. None. 1969. Type: C. asclepiadeum (Willd.) Fr. 1815, basionym: Type: E. harknessii (J.P. Moore) Y. Hirats. 1969, Erineum asclepiadeum Willd. 1806, now regarded basionym: Peridermium harknessii J.P. Moore 1884, as Cronartium flaccidum (Alb. & Schwein.) G. Winter now regarded as Cronartium harknessii (J.P. Moore) 1880. E. Meinecke 1920. Elateraecium Thirum. et al. in Mycologia 58: 394. Hiratsukamyces Thirum. et al. Sydowia 27: 80, 1975. None. 1966. Type: E. salaciicola Thirum. et al. 1966. Type: H. salaciicola Thirum. et al. 1975, now regarded as Elateraecium salaciicola Thirum. et al. 1966. Gerwasia Racib. in Bull. Int. Acad. Sci. Cracovie, Cl. Morispora Salazar-Yepes et al. in Caldasia 29: 108. None. Sci. Math. 3: 270. 1909. 2007. Type: G. rubi Racib. 1909. Type: M. tenella (H.S. Jacks. & Holw.) Salazar-Yepes et al. 2007, basionym: Mainsia tenella H.S. Jacks. & Holw. 1931, now regarded as Gerwasia tenella (H.S. Jacks. & Holw.) Buriticá 1994. Gymnosporangium R. Hedw. ex DC., in Lamarck & Roestelia Rebent., Prodr. fl. neomarch.: 350. 1804. Protection needed by NCF. de Candolle, Fl. franç., ed. 3 2: 216. 1805. Type: G. fuscum DC. 1805, now regarded as Type: R. cancellata (Pers.) Rebent. 1804, basionym: Gymnosporangium sabinae (Dicks.) G. Winter 1884. Aecidium cancellatum Pers. 1792, now regarded as Gymnosporangium sabinae (Dicks.) G. Winter 1884. Helicobasidium Pat. in Bull. Soc. Bot. France 32: Thanatophytum Nees, Syst. Pilze: 148. 1816. Protection needed by NCF. 172. 1885. Type: H. purpureum (Tul.) Pat. 1885, basionym: Type: T. crocorum (Pers.) Nees 1816, basionym: Hypochnus purpureus Tul. 1865. Sclerotium crocorum Pers. 1801, now regarded as Helicobasidium purpureum (Tul.) Pat. 1885. Tuberculina Tode ex Sacc. in Michelia 2: 34. 1881. Type: T. persicina (Ditmar) Sacc. 1881, basionym: Tubercularia persicina Ditmar 1817, now regarded as Helicobasidium purpureum (Tul.) Pat. 1885. Hyalopycnis Höhn. in Hedwigia 60: 152. 1918. Heterogastridium Oberw. & R. Bauer in Mycologia 82: None. 57. 1990. Type: H. hyalina Höhn. 1918, now regarded as Type: H. pycnidioideum Oberw. & R. Bauer 1990. Heterogastridium pycnidioideum Oberw. & R. Bauer 1990.

VOLUME 9 · NO. 1 77 Aime et al.

Table 1. (Continued).

Recommended generic name Synonymous alternate morph generic name(s) Action required

Kriegeria Bres. in Rev. Mycol. (Toulouse) 13: 14. Xenogloea Syd. & P. Syd. in Ann. Mycol. 17: 44. None. 1891. 1919. ARTICLE Type: K. eriophori Bres. 1891. Type: X. eriophori (Bres.) Syd. & P. Syd. 1919, basionym: Kriegeria eriophori Bres. 1891. Zymoxenogloea D.J. McLaughlin & Doublés in Mycologia 84: 671. 1992. Type: Z. eriophori D.J. McLaughlin & Doublés 1992, now regarded as Kriegeria eriophori Bres. 1891. Kuehneola Magnus in Bot. Centralbl. 74: 169. 1898. Spirechina Arthur in J. Mycol. 13: 30. 1907. None. Type: K. albida (J.G. Kühn) Magnus 1898, basionym: Type: S. loeseneriana (Henn.) Arthur 1907, basionym: Chrysomyxa albida J.G. Kühn 1883, now regarded as Uredo loeseneriana Henn. 1898, now regarded as Kuehneola uredinis (Link) Arthur 1906. Kuehneola loeseneriana (Henn.) H.S. Jacks. & Holw. 1931. Melampsorella J. Schröt. in Hedwigia 13: 85. 1874. Peridermium (Link) J.C. Schmidt & Kunze in Protection needed by NCF. Deutschl. Schwämme 6: 4. 1817, basionym: Hypodermium subgen. Peridermium Link 1816. Type: M. caryophyllacearum (DC.) J. Schröt. 1874, Type: Aecidium elatinum Alb. & Schwein. 1805, now basionym: Uredo caryophyllacearum DC. 1805. regarded as Melampsorella caryophyllacearum (DC.) J. Schröt. 1874. Milesina Magnus in Ber. Deutsch. Bot. Ges. 27: 325. Milesia F.B. White in Scott. Nat. 4: 162. 1878. Protection needed by NCF. 1909. Type: M. kriegeriana (Magnus) Magnus 1909, Type: M. polypodii F.B. White 1878, now regarded as basionym: Melampsorella kriegeriana Magnus 1901. Milesina polypodii (F.B. White) Aime & Rossman 2018. Phragmidium Link in Ges. Naturf. Freunde Berlin Aregma Fr., Observ. Mycol. 1: 22., 1815. Protection needed by NCF. Mag. 7: 30. 1816. Type: P. mucronatum (Pers.) Schltdl. 1824, basionym: Type: A. obtusatum Fr. 1832, now regarded as Puccinia mucronata Pers. 1794. Phragmidium fragariae (DC.) G. Winter 1884, basionym: Puccinia fragariae DC., 1808. Lecythea Lév. in Ann. Sci. Nat., Bot., sér. 3 8: 373. 1847. Type: L. miniata (Pers.) Lév. 1847, basionym: Uredo miniata Pers. 1801, now regarded as Phragmidium mucronata (Pers.) Schltdl. 1824. Prospodium Arthur in J. Mycol. 13: 31. 1907. Canasta A.A. Carvalho & J.F. Hennen in Mycologia None. 102: 1096. 2010. Type: P. appendiculatum (G. Winter) Arthur 1907, Type: C. cruscula A.A. Carvalho & J.F. Hennen 2010, basionym: Dicaeoma appendiculata Kuntze 1898. now regarded as Prospodium cruscula (A.A. Carvalho & J.F. Hennen) Aime & Rossman 2018. Puccinia Pers. in Neues Mag. Bot. 1: 118. 1794. Aecidium Pers., Observ. Mycol. 1: 97. 1796. None. Type: P. graminis Pers. 1794. Type: A. berberidis Pers. 1796, now regarded as Puccinia graminis Pers. 1794. Caeoma Link in Ges. Naturf. Freunde Berlin Mag. 3: 5. 1809. Type: C. berberidis (Pers.) Har. 1809, basionym: Aecidium berberidis Pers. 1792, now regarded as Puccinia graminis Pers. 1794. Rhodotorula F.C. Harrison in Proc. & Trans. Roy. Chromotorula F.C. Harrison in Proc. & Trans. Roy. None. Soc. Canada, ser. 3 21: 349. 1927. Soc. Canada, ser. 3 21: 350. 1927. Type: R. glutinis (Fresen.) F.C. Harrison 1928. Type: C. kitae F.C. Harrison 1927. Rhodosporidium Banno in J. Gen .Appl. Microbiol. 13: 192. 1967. Type: R. toruloides Banno 1967, now regarded as Rhodotorula toruloides (Banno) Q.M. Wang et al. 2015. Septobasidium Pat. in J. Bot. (Morot) 6: 63. 1892. Johncouchia S. Hughes & Cavalc. in Canad. J. Bot. None. 61: 2226. 1983. Type: S. velutinum Pat. 1892. Type: J. mangiferae (Bat.) S. Hughes & Cavalc. 1983, basionym: Cladotrichum mangiferae Bat. 1956, now regarded as Septobasidium pilosum Boedijn & B.A. Steinm. 1930.

78 IMA FUNGUS Asexual-sexual generic names in Pucciniomycotina and Ustilaginomycotina

Table 1. (Continued). ARTICLE

Recommended generic name Synonymous alternate morph generic name(s) Action required

Sporobolomyces Kluyver & C.B. Niel in Centralbl. Aessosporon Van der Walt in Antonie van Protection needed by NCF. Bakteriol., 2 Abt. 2: 63. 1924. Leeuwenhoek 36: 54. 1970.

Type: S. salmonicolor (B. Fisch. & Brebeck) Kluyver & Type: A. salmonicolor Van der Walt 1970, now C.B. Niel 1924, basionym: Blastoderma salmonicolor regarded as Sporobolomyces salmonicolor (B. Fisch. B. Fisch. & Brebeck 1894. & Brebeck) Kluyver & C.B. Niel 1924. Blastoderma B. Fisch. & Brebeck, in Fischer, Morph. Syst.: 47. 1894. Type: B. salmonicolor B. Fisch. & Brebeck 1894, now regarded as Sporobolomyces salmonicolor (B. Fisch. & Brebeck) Kluyver & C.B. Niel 1924. Rhodomyces Wettst. in Sitzungsber. Kaiserl. Akad. Wiss., Wien, Math.-naturw. Kl., Abt. 1 91: 39. 1885. Type: R. kochii Wettst. 1885, now regarded as Sporobolomyces salmonicolor (B. Fisch. & Brebeck) Kluyver & C.B. Niel 1924. Sporidiobolus Nyland in Mycologia 41: 686. 1950. Type: S. johnsonii Nyland 1950, now regarded as Sporobolomyces johnsonii (Nyland) Q.M. Wang et al. 2015. Uromyces (Link) Unger, Exanth. Pflanzen: 277. Uredo Pers., Syn. Meth. Fung. 1: 214. 1801. Protection needed by NCF. 1833 (nom. cons.), basionym: Hypodermium subgen. Uromyces Link 1816. Type: U. appendiculata (Pers.) Unger 1796, basionym: Type: U. betae Pers. 1801, now regarded as Uredo appendiculatus Pers., now regarded as: Uromyces beticola (Bellynck) Boerema, Loer. & Uromyces appendiculatus (Pers.) Link 1816. Hamers 1987.

as sexual and asexual morphs of the same species and Blastospora = 567, Pelastoma = 11), we recommend the clarified the terminology associated with them, thus these use of Blastospora for this genus. generic names are synonyms. Given that Basidiopycnis has priority and both generic names are monotypic and Two new combinations are required: relatively obscure (GS Basidiopycnis = 37, Basidiopycnides = 5), we follow the principle of priority and recommend the Blastospora juruensis (Henn.) Aime & Rossman, use of Basidiopycnis. comb. nov. MycoBank MB824643 Use Blastospora Dietel 1908 (S) rather than Basionym: Uredo juruensis Henn., Hedwigia 43: 161 (1904). Pelastoma Salazar-Yepes et al. 2012 (A) Synonyms: Hemileia juruensis (Henn.) P. Syd. & Syd., Mon. Ured. 3: 219 (1915). The generic name Blastospora typified by B. smilacis Hemileia jahnii Syd., Ann. Mycol. 28: 46 (1930). includes tropical heteroecious rust fungi as summarized Uredo jahnii (Syd.) Ritschel, Biblioth. Mycol. 200: 98 (2005). by Salazar-Yepes et al. (2012) who described Pelostoma Hemileia colombiana Buriticá, Caldasia 12: 165 (1978). typified byP. juruense for the asexual morph of B. columbiana Blastospora colombiana (Buriticá) Salazar-Yepes et al., (syn. Hemileia columbiana). Aime (2006) demonstrated that Mycologia 104: 116 (2012). B. smilicis has affinities to Hemileia and Maravalia in the Mikronegeriaceae. Blastospora smilacis and B. itoana were Blastospora subneurophyla (Speg.) Aime & considered to have asexual morphs belonging to Caeoma Rossman, comb. nov. (Ono et al. 1986, 1987); however, the type of Caeoma is C. MycoBank MB824644 berberidis, a synonym of Puccinia graminis, thus Caeoma is Basionym: Uredo subneurophyla Speg., Anales Soc. Ci. a synonym of Puccinia. No other species of Blastospora nor Argent. 17: 123 (1883). the second species of Pelostoma have been sequenced. Synonyms: Puccinia subneurophyla (Speg.) Speg., Anales Assuming that B. columbiana is congeneric with B. smilicis, Mus. Nac. Hist. Nat. Buenos Aires 31: 384 (1922). then Blastospora and Pelostoma are synonyms. At present Pelastoma subneurophylum (Speg.) Salazar-Yepes et al., five species are included in Blastospora and one name Mycologia 104: 116 (2012). additional to the type has been placed in Pelostoma. Uredo forsteroniae Henn., Hedwigia 34: 99 (1895). Considering the fact that Blastospora has priority, a greater number of names, and is more commonly used (GS

VOLUME 9 · NO. 1 79 Aime et al.

Table 2. Recommended generic names of Ustilaginomycotina among sexual-asexual names that compete for use. The recommended name is in bold; see text for the rationale behind these recommendations. For each generic name this list provides the author, its date and place of publication, type species of the genus, its basionym, their dates of publication and the currently accepted name, if different. The action required is indicated in the last column such as approval by the Nomenclature Committee for Fungi (NCF) for those generic names that do not have priority and thus need protection.

Recommended generic name Synonymous alternate morph generic name(s) Action required ARTICLE

Anthracoidea Bref., Unters. Gesammtgeb. Mykol. 12: 143. Crotalia Liro in Ann. Acad. Sci. Fenn., Ser. A 42(1): 545, None. 1895. 273. 1938. Type: A. caricis (Pers.) Bref. 1896, basionym: Uredo caricis Type: C. cintractiae-fischeri Liro 1939, now regarded as Pers. 1801. Anthracoidea fischeri (P. Karst.) Kukkonen 1963. Doassansia Cornu in Ann. Sci. Nat., Bot., sér. 6 15: 285. Savulescuella Cif. in Bontea et al. [eds], Omag. None. 1883. Savulescu: 179. 1959. Type: D. alismatis (Nees ex Fr.) Cornu 1883, basionym: Type: S. alismacearum (Sacc.) Cif. 1959, basionym: Sclerotium alismatis Nees ex Fr. 1822. Cylindrosporium alismacearum Sacc. 1880, now regarded as Doassansia alismatis (Nees ex Fr.) Cornu 1883.

Doassansiopsis (Setch.) Dietel in Engler & Prantl, Nat. Doassansiella Zambett. in Rev. Mycol. (Paris) 35: 164. None. Pflanzenfam.1 (1**): 15. 1897, basionym: Doassansia 1970. subgen. Doassansiopsis Setch. 1891. Type: Doassansia deformans Setch. 1892, now regarded as Type: D. aquatilis (Peck) Zambett. 1970, basionym: Doassansiopsis deformans (Setch.) Dietel 1897. Ramularia aquatilis Peck 1884, now regarded as Doassansiopsis hydrophila (A. Dietr.) Lavrov 1937. Entyloma de Bary in Bot. Zeitung (Berlin) 32(7): 101, 1874. Entylomella Höhn. in Ann. Mycol. 22: 191. 1924. None. Type: E. microsporum J. Schröt. 1874. Although some Type: E. ranunculi (Bonord.) Höhn. 1916, basionym: have attributed E. microsporum to Protomyces microsporus Fusidium ranunculi Bonord. 1851, now regarded as F. Unger 1834, this name is a later homonym of P. Entyloma ficariae F. Thümen ex A.A. Fischer v. Waldheim microsporus Pass. 1833. 1877. Mycosarcoma Bref., Unters. Gesammtgeb. Mykol. 15: 53. Pseudozyma Bandoni in J. Linn. Soc., Bot. 91: 38. 1985. Awaiting decision 1912. of NCF on the proposal of Type: P. prolifica Bandoni 1985, now regarded as Type: M. maydis (DC.) Bref. 1912, basionym: Uredo maydis Thines (2016) Mycosarcoma maydis (DC.) Bref. 1912 or Ustilago DC. 1815. as to whether maydis (DC.) Corda 1842. Mycosarcoma OR should be taken Ustilago (Pers.) Roussel, Fl. Calvados, ed. 2: 4. 1806, up. basionym: Uredo [unranked] Ustilago Pers. 1801. Type: Ustilago hordei (Pers.) Lagerh. 1889, basionym: Uredo segetum a hordei Pers. 1801, or with a newly conserved type proposed by Thines (2016): Ustilago maydis (DC.) Corda 1842, basionym: Uredo maydis DC. 1815.

Thecaphora Fingerh. in Linnaea 10: 230. 1836. Angiosorus Thirum. & M.J. O’Brien 1974 in Sydowia 26: None. 201 (1974) [1972]. Type: T. hyalina Fingerh. 1836, now regarded as Type: A. solani Thirum. & M.J. O’Brien 1974, now Thecaphora seminis-convolvuli (Desm.) S. Ito 1935. regarded as Thecophora solani (Thirum. & M.J. O’Brien) Mordue 1988. Rhombiella Liro in Ann. Acad. Sci. Fenn., Ser. A 42(1): 507, 545. 1939. Type: R. cardamines Liro 1939, now regarded as Thecaphora thlaspeos (Beck) Vánky 2004. Thecaphorella H. Scholz & I. Scholz in Nova Hedwigia 47: 71. 1988. Type: T. antherarum (Oudem.) H. Scholz & I. Scholz 1988, basionym: Gloeosporium antherarum Oudem. 1898, now regarded as Thecaphora seminis-convolvuli (Desm.) S. Ito 1935. Tilletia Tul. & C. Tul. in Ann. Sci. Nat., Bot., sér. 3 7: 112. Tilletiella Zambett. in Rev. Mycol. (Paris) 35: 165. 1970. None. 1847. Type: Tilletia caries (DC.) Tul. & C. Tul. 1847, basionym: Type: T. alopecuri (Sawada) Zambett. 1970, basionym: Uredo caries DC. 1815, now regarded as T. tritici (Bjerk.) G. Entyloma alopecuri Sawada 1922, now regarded as Winter 1875, basionym: Lycoperdon tritici Bjerk. 1775. Tilletia alopecuri (Sawada) L. Ling 1949.

80 IMA FUNGUS Asexual-sexual generic names in Pucciniomycotina and Ustilaginomycotina

Table 2. (Continued). ARTICLE Recommended generic name Synonymous alternate morph generic name(s) Action required Urocystis Rabenh. ex Fuckel in Jahrb. Nassauischen Paepalopsis J.G. Kühn in Irmischia 7: 39, 1882. None. Vereins Naturk. 23–24: 41. 1870 (nom. cons.). Type: U. occulta (Wallr.) A.A. Fisch. Waldh. 1867, basionym: Type: P. irmischiae J.G. Kühn 1882, now regarded as Erysibe occulta Wallr. 1833. Urocystis primulae (Rostr.) Vánky 1985. Polycystis Lév. in Ann. Sci. Nat. Bot., Ser. 3, 5: 269. 1846. Type: P. pompholygodes Lév. 1846, now regarded as Urocystis anemones (Pers.) G. Winter 1880.

Use Camptobasidium Marvanová & Suberkr. All species of Endocronartium have been shown to 1990 (S) rather than Crucella Marvanová & belong in the genus Cronartium either as synonyms or Suberkr. 1990 (A) distinct species within that genus. Several new combinations are made below. In addition, Endocronartium pini (Willd.) Y. The generic names Camptobasidium and Crucella were Hirats. (syn. Peridermium pini (Willd.) Lév.) has been shown described in the same article based on sexually and to be a synonym of C. flaccidum (Hantuka et al. 2002). asexually typified morphs of the same species, thus they Although some of the asexual morphs of species of are synonyms and compete equally for use (Marvanová &. Cronartium have been recognized in Peridermium, that generic Suberkropp 1990). Both are monotypic and have been used name is now regarded as a synonym of Melampsorella based to the same extent, although Crucella is also the name of a on the type species, Aecidium elatinum, which is a later name fossil radiolarian, thus it is difficult to determine its usage as for M. caryophyllacearum (see below under Melampsorella). a fungus. Given the confusion about the use of the name Thus, names in Peridermium must be recognized in the Crucella and because preference has previously been sexually typified genus in which the species belong, primarily given to sexual morph names, we recommend the use of Cronartium but also Chrysomyxa and Coleosporium. Camptobasidium. The following new combinations are required:

Protect Classicula R. Bauer et al. 2003 (S) over Cronartium bethelii (Hedgc. & Long) Aime & Naiadella Marvanová & Bandoni 1987 (A) Rossman, comb. nov. MycoBank MB824645 The monotypic generic name Classicula is typified by C. Basionym: Peridermium bethelii Hedgc. & Long, fluitans, which was described as the sexual morph of N. Phytopathology 3: 251 (1913). fluitans (Bauer et al. 2003), the type species of the monotypic generic name Naiadella, thus these names are synonyms. Vogler & Bruns (1998) demonstrated that Peridermium Neither name is frequently used (GS Classicula = 73; bethelii belongs in Cronartium, as sister to C. comandrae. Naiadella = 86); however, the family Classiculaceae, order With the change in the ICN, it is possible to place this name Classiculales, and class Classiculomycetes are based on the in Cronartium. name Classicula, thus we recommend the use of Classicula. Cronartium kurilense (Dietel) Y. Ono, comb. nov. Use Cronartium Fr. 1815 (S) rather than MycoBank MB824647 Endocronartium Y. Hirats. 1969 (S) Basionym: Peridermium kurilense Dietel, Bot. Jahrb. Syst. 37: 107 (1905), The generic name Cronartium is well known for species that Synonym: Endocronartium sahoanum var. hokkaidoense cause stem rusts on conifers in temperate regions, often Imazu & Kakish., Trans. Mycol. Soc. Japan 33: 174 with alternate hosts on herbaceous plants. One of the most (1992). notorious species is C. flaccidum, cause of Scots pine blister rust, of which C. asclepiadeum, the type species of Cronartium, A blister rust fungus on Pinus pumila in the North Kuril is a synonym (Mordue & Gibson 1978). Endocronartium was Islands (southeast of Kamchatka, Russia) was described as established for species genetically similar to Cronartium Peridermium kurilense, distinct from C. ribicola (Dietel 1905). except for being autoecious (i.e. completing their life-cycle It was subsequently synonymized under C. kamtschaticum on a single host) and endocyclic (lacking aecia and uredinia (Jørstad 1934, Hiratsuka et al. 1992). An endocyclic fungus and having aecioid telia) (Hiratsuka 1969). The type species on Pinus pumila distributed in northern Honshu, Japan, was of Endocronartium, E. harknessii, cause of western gall rust, named as E. sahoanum (Imazu et al. 1989), and a similar has long been known to be indistinguishable from species fungus on the same host species distributed in Hokkaido of Cronartium except for the life-cycle reductions (Epstein & was separated as a variety, E. sahoanum var. hokkaidoense, Buurlage 1988). With the change in the ICN, these generic distinct from E. sahoanum var. sahoanum on the basis of names are synonyms and the six taxa of Endocronartium differences in spore and germ tube morphology (Imazu & should be recognized in the older, commonly used genus Kakishima 1992). Based on the similarity in spore shape, Cronartium. size, surface structure, and PCR-RFLP pattern, P. kurilense

VOLUME 9 · NO. 1 81 Aime et al.

and E. sahoanum var. hokkaidoense have been shown to be Kern 1973), thus these generic names are synonyms. Although conspecific and distinct fromC. kamtschaticum, which is now 55 names have been placed in Roestelia, their sexual morphs considered a synonym of C. ribicola (Imazu et al. 2000, Kim belong primarily in Gymnosporangium. The need to protect et al. 2010). Thus, P. kurilense must be placed in Cronartium. Gymnosporangium was recognized many years ago; however, the proposal to conserve Gymnosporangium over Roestelia

ARTICLE Cronartium sahoanum (Imazu & Kakish.) Aime & was rejected for lack of adequate justification (Rogers 1953). Rossman, comb. nov. Because Gymnosporangium is by far the most commonly used MycoBank MB824648 name rather than Roestelia (GS Gymnosporangium = 6050, Basionym: Endocronartium sahoanum Imazu & Kakish. Roestelia = 598) and includes a greater number of species, we Trans. Mycol. Soc. Japan 30: 308 (1989). recommend the protection of Gymnosporangium.

Cronartium yamabense (Saho & I. Takah.) Aime & Protect Helicobasidium Pat. 1885 (S) over Rossman, comb. nov. Thanatophytum Nees 1816 (A) and Tuberculina MycoBank MB824649 Tode ex Sacc. 1880 (A) Basionym: Peridermium yamabense Saho & I. Takah., Trans. Mycol. Soc. Japan 22: 33 (1981). Helicobasidium, typified by H. purpureum, includes 15 species, Synonym: Endocronartium yamabense (Saho & I. Takah.) some of which cause plant diseases, such as H. purpureum, the Paclt, Mitt. Deutsch. Dendrol. Ges. 77: 227 (1987). cause of cosmopolitan violet root rot, and the closely related H. mompa, also causing violet root rot in Africa and Asia (Sayama Use Elateraecium Thirum. et al. 1966 (A) rather et al. 1994). The asexual morph of H. purpureum has been than Hiratsukamyces Thirum. et al. 1975 (S) regarded as Thanatophytum crocorum (Roberts 1999). Two species were mentioned in the protolog for Thanatophytum; Donk The generic name Hiratsukamyces, typified by H. salaciicola, (1962) lectotypified this genus with T. crocorum. The generic was established for the sexual morph of Elateraecium name Tuberculina is lecto-typified by T. persicina (Clements & salaciicola, type species of Elateraecium (Thirumalachar et Shear 1931), a parasite of rust fungi. The relationship between al. 1966, 1975), thus these generic names are synonyms. these names all representing the same species was confirmed Two names were included in Hiratsukamyces when it was by Lutz et al. (2004) in which the life strategy of this unusual described, each also having a name in Elateraecium for the fungus was clarified. More specifically, they determined that asexual morph. Since then two more names have been added T. persicina is a synonym of Helicobasidium purpureum. The to Elateraecium by Gjaerum & Reid (1983). Both generic commonly used name H. purpureum based on Hypochnus names are obscure but given the greater number of names, purpureus is herein proposed for protection over the older names priority, and slightly greater use (GS Elateraecium = 12, Sclerotium crocorum and Tubercularia persicina. Among these Hiratsukamyces = 11), we recommend the use of Elateraecium. three generic names, Helicobasidium is most commonly used (GS Helicobasidium = 2090, Thanatophytum = 67, Tuberculina Use Gerwasia Racib. 1909 (S) rather than = 487). Given the widespread use of Helicobasidium and the Morispora Salazar-Yepes et al. 2007 (A) number of species in this genus, we recommend Helicobasidium for protection. In addition we recommend the name of the type Gerwasia, typified by G. rubi, is a well-known genus of species, H. purpureum, for protection. autoecious rust fungi that occurs on Rosaceae, primarily in the Asian and American tropics (Cummins & Hiratsuka Name for protection: 2003). Twenty-two species have been described in Gerwasia of which 12 have asexual morphs that have been placed in Helicobasidium purpureum (Tul.) Pat., Bull. Soc. Bot. Morispora, typified by M. tenella, a species with a sexual France 32: 171 (1885). morph regarded as G. tenella (Salazar-Yepes et al. 2007). Basionym: Hypochnus purpureus Tul., Ann. Sci. Nat., Bot., Thus, assuming that G. rubi and G. tenella are congeneric sér. 5 4: 295 (1865); nom. prot. prop. as indicated by Salazar-Yepes et al. (2007) but without Rejected names and synonyms: Tuber parasiticum Bull., molecular data, Gerwasia and Morispora are synonyms. Hist. champ. France 1: 81 (1791). All names in Morispora already have a name in Gerwasia. Sclerotium crocorum Pers., Syn. meth. fung. 1: 119 (1801). Given its priority, greater number of names, and widespread Rhizoctonia crocorum (Pers.) DC., in De Candolle & Lamarck, use (GS Gerwasia = 112, Morispora = 6), the generic name Fl. franç., 3rd edn 5: 110 (1815). Gerwasia is recommended for use. Thanatophytum crocorum (Pers.) Nees, Syst. Pilze: 148 (1816) [“1816-17”]. Protect Gymnosporangium R. Hedw. ex DC. Rhizoctonia medicaginis DC., in De Candolle & Lamarck, Fl. 1805 (S) over Roestelia Rebent. 1804 (A) franç., 3rd edn 5: 111 (1815). Tubercularia persicina Ditmar, in Sturm, Deutschl. Fl., Abt. 3 The genus Gymnosporangium, based on G. fuscum, now 1: 99 (1817). recognized as G. sabinae, includes ubiquitous rust fungi that Tuberculina persicina (Ditmar) Sacc., Fungi italice autogr. have been well-studied (Yun et al. 2009). Index Fungorum lists del. 17-28: tab. 964 (1881). 145 names in this genus. The type species of Roestelia, R. Protonema brebissonii Desm., Pl. crypt. N. France: no. 651 cancellata, is a synonym of G. sabinae (Alexopoulos 1940, (1834).

82 IMA FUNGUS Asexual-sexual generic names in Pucciniomycotina and Ustilaginomycotina

Helicobasidium brebissonii (Desm.) Donk, Taxon : 164 been used for the asexual morph of species of Kuehneola 7 ARTICLE (1958). and Gerwasia. Spirechina loeseneriana is regarded as the Rhizoctonia rubiae M.J. Decne., Rech. Anat. Physiol. asexual morph of K. loeseneriana (Jackson 1931). Although Garance: 55 (1837). 13 species had been named in Spirechina, all have been Rhizoctonia violacea Tul. & C. Tul., Fung. hypog.: 188 (1851). placed in Gerwasia, Kuehneola, or Mainsia (Kirk 2017). Given its priority, greater number of names, and more Among the extensive list of synonymous names, frequent use (GS Kuehneola = 590, Spirechina = 42), we Helicobasidium purpureum is the name in Helicobasidium recommend Kuehneola for use. most commonly used for this important and ubiquitous plant pathogen (GS Helicosporium purpureum = 627, H. Protect Melampsorella J. Schröt. 1874 (S) over berbissonii = 7). Although several names for this fungus have Peridermium (Link) J.C. Schmidt & Kunze 1817 been placed in Rhizoctonia, the type species of Rhizoctonia (A) is R. solani, which represents a genus in Agaricomycotina. The type species of Melampsorella is M. caryophyllacearum, Use Hyalopycnis Höhn. 1918 (A) rather than while Peridermium is typified with the conserved type, Heterogastridium Oberw. & R. Bauer 1990 (S) Aecidium elatinum, as formally proposed by Hiratsuka (1974) and approved (Petersen 1975). Because the latter is regarded The generic name Hyalopycnis is typified by H. hyalina. as a synonym of M. caryophyllacearum (Hiratsuka 1974), Hyalopycnis blepharistoma, a synonym of H. hyalina, these generic names are synonyms. Melampsorella has five was regarded as the asexual morph of Heterogastridium names in Index Fungorum. The asexually typified generic pycnidioidium, the type of the monotypic generic name name Peridermium includes many species most of which Heterogastridium as explained by Oberwinkler et al. (1990) have sexual morphs in Cronartium (Vogler & Bruns 1998). in describing the order Heterogastridiales, now placed Given the relationship of species of Peridermium with both in Microbotryomycetes, Pucciniomycotina (Aime et al. Melampsorella and Cronartium, we recommend abandoning 2006, Toome & Aime 2014). Thus, the generic names the name Peridermium in favour of Melampsorella. Heterogastridium and Hyalopycnis are synonyms. Although Although the type species of Peridermium dictates the generic name Heterogastridium has been used slightly that this generic name is a synonym of Melampsorella, more frequently (GS Heterogastridium = 54, Hyalopycnis = a number of names introduced in Peridermium belong in 43) and serves as the basis for the ordinal name, Hyalopycnis other rust genera, such as Cronartium, Coleosporium and has priority and its use circumvents the need for a name several genera of Pucciniastraceae. An effort will be made change for the only species, thus we recommend the use of to determine the correct generic placement of species Hyalopycnis. Its use does not affect the ordinal name. described in Peridermium. Furthermore, an asexual morph can still be referred to as peridermium-like, using the term in Use Kriegeria Bres. 1891 (S) rather than a non-latinized form to describe a morphology. Xenogloea Syd. & P. Syd. 1919 (S) and Zymoxenogloea D.J. McLaughlin & Doublés Protect Milesina Magnus 1909 (S) over Milesia 1992 (A) F.B. White 1878 (A) The monotypic generic name Kriegeria Bres., typified by The genus Milesina includes over 60 species, primarily K. eriophori, was determined to be legitimate by Toome et occurring on Abies alternating with ferns in several families. al. (2015). Previously it has been confused with Kriegeria The type species, Milesina kriegeriana, occurs on Abies Höhnel 1914 and Kriegeria Rabenh. ex Seaver 1943, whose and Dryopteris in Europe and is known to have an asexual species are now placed in other genera in the ascomycete morph recognized as Milesia kriegeriana. The type species families Helotiaceae and Rutstroemiaceae. Kriegeria Bres. of Milesia is Milesia polypodii, recognized as the uredinial 1891 has also been confused with the later monotypic state of Milesina dieteliana (Laundon 1967), thus Milesina generic name Xenogloea typified by the same species, X. and Milesia are synonyms. Although Milesia is the older eriophori. An asexual morph of K. eriophori, as X. eriophori, generic name and has about the same number of names as was named Zymoxenogloea eriophori, monotype species Milesina (72 vs. 67, respectively), Milesina is in broader use of Zymoxenogloea, thus Kriegeria Bres. 1891, Xenogloea, especially in recent literature (e.g. Berndt 2007, Kakishima and Zymoxenogloea are synonyms. Following Toome et al. et al. 2016, Klenke & Scholler 2015, Smith 2015, Tanese & (2015) and priority, we recommend the use of Kriegeria. Negrean 2007, Wingfield et al. 2004). For these reasons, we recommend the protection and use of Milesina. Use Kuehneola Magnus 1899 (S) rather than Spirechina Arthur 1907 (A) One new combination is required:

The generic name Kuehneola, typified by K. albida, now Milesina polypodii (F.B. White) Aime & Rossman, regarded as K. uredinis (Ono 2015), includes about 25 comb. nov. species of rust fungi that occur primarily on species of Rubus MycoBank MB824650 and other Rosaceae (Cummins & Hiratsuka 2003). The Basionym: Milesia polypodii F.B. White, Scott. Naturaist generic name Spirechina, typified by S. loeseneriana, has (Perth). 4: 163 (1878) [“1877-1878”]

VOLUME 9 · NO. 1 83 Aime et al.

Synonyms: Milesina dieteliana (Syd. & P. Syd.) Magnus, Ber. this major food source throughout history (Schumann 1991). Deutsch. Bot. Ges. 27: 325 (1909). Several thousand names have been placed in Puccinia. Melampsorella dieteliana Syd. & P. Syd., Ann. Mycol. 1(6): The asexually typified generic name Aecidium is typified by 537 (1903). A. berberidis, a name used for one of the alternate morphs of P. graminis (Cummins 1971), thus these generic names

ARTICLE Protect Phragmidium Link 1816 (S) over are synonyms. Prior to 2012, Aecidium was used when a Aregma Fr. 1815 (S) and use rather than sexual morph was lacking for rust fungi that appeared to Lecythea Lév. 1847 (A) have affinities with Puccinia. Before the changes in the ICN, the species could not be described in the “correct” genus The generic name Phragmidium is applied to a group of rust (Puccinia) even though it was possible to determine its species that occur on Rosaceae. Recent molecular studies generic placement. The need for the generic name Aecidium have shown that Phragmidium is a well-defined genus in no longer exists. The type species of Caeoma, C. berberidis, Phragmidiaceae and that the type species of Phragmidium, is a synonym of Puccinia graminis (Laundon 1967), thus P. mucronatum, is distinct from the other species commonly Caeoma is a synonym of Puccinia. Given the ubiquitous use encountered on dog rose, P. tuberculatum (Ritz et al. 2005, of the generic name Puccinia, it is recommended for use. Yun et al. 2011). Both those papers also demonstrated that P. mucronatum is congeneric but distinct from P. fragariae, the Use Rhodotorula F.C. Harrison 1928 (A) rather correct name for the type species of Aregma, A. obtusatum; than Chromotorula F.C. Harrison 1928 (A) and thus, Aregma and Phragmidium are synonyms. Several Rhodosporidium Banno 1967 (S) species are mentioned in the protolog for Aregma; this genus was lectotypified by Laundon (1965). The type species of Rhodotorula, R. glutinis, was shown The asexually typified generic name Lecythea is typified to be congeneric with Rhodosporidium toruloides, type of by L. miniata, based on Uredo miniata, and now regarded as Rhodosporidium, by Wang et al. (2015a). They recognized a synonym of P. mucronatum. Although Lecythea includes 27 the priority of Rhodotorula and placed Rhodosporidium names, all but four have been placed in other genera (see Index toruloides in Rhodotorula. All 14 names described in Fungorum). Most of the 16 names in Aregma have been placed Rhodosporidium have been placed in two genera, or have synonyms in Phragmidium, while 250 names have Rhodosporidiobolus and Rhodotorula. More than 150 names been described in Phragmidium; this generic name is much have been introduced in Rhodotorula. Chromotorula, typified more commonly used than Aregma (GS Phragmidium = 3770, by C. kitae, was published in the same article as Rhodotorula Aregma = 244). Given its common usage and greater number of and thus has equal priority to Rhodotorula. Chromotorula has names, we recommend that Phragmidium be protected. long been considered a synonym of Rhodotorula (Statzell- Tallman & Fell 1998), and all but two of the six names in Use Prospodium Arthur 1907 (S) rather than Chromotorula have been placed elsewhere. In agreement Canasta A.A. Carvalho & J.F. Hennen 2010 (A) with Wang et al. (2015a) as well as priority, greater number of names, and most frequent use (GS Rhodotorula = 50 300, The generic name Prospodium includes about 50 species Rhodosporidium = 6130 Chromotorula = 30), we recommend occurring on Bignoniaceae and Verbenaceae in tropical the use of Rhodotorula. regions (Cummins & Hiratsuka 2003, Hernández & Hennen 2003). The generic name Canasta was established for Use Septobasidium Pat. 1892 (S) rather than asexual morphs of species of Prospodium (Carvalho & Johncouchia S. Hughes & Cavalc. 1983 (A) Hennen 2010), although the type species C. cruscula does not have a known sexual morph. Given that the other two The monotypic generic name Johncouchia, typified by names described in Canasta have sexual morphs that are J. mangiferae, was described for the asexual morph of species of Prospodium, it seems likely that Canasta is a Septobasidium pilosum (Hughes & Cavalcanti 1983). synonym of Prospodium. Given its priority, greater number Septobasidium pilosum and S. velutinum, the type species of species and widespread use, we recommend the use of of Septobasidium, were determined to be congeneric by Prospodium. One new combination is necessary: Henk & Vilgalys (2007). Septobasidium includes over 300 names, has priority and is widely used (GS Septobasidium = Prospodium crusculum (A.A. Carvalho & J.F. Hennen) 1200, Johncouchia = 15), and so we recommend the use of Aime & Rossman, comb. nov. Septobasidium. MycoBank MB824651 Septobasidium has already been formally conserved Basionym: Canasta cruscula A.A. Carvalho & J.F. Hennen, against Campylobasidium Lagerh. ex F. Ludw. 1892, Mycologia 102: 1098 (2010) Gausapia Fr. 1825, and Glenospora 1849.

Use Puccinia Pers. 1794 (S) rather than Protect Sporobolomyces Kluyver & C.B. Niel Aecidium Pers. 1796 (A) and Caeoma Link 1924 (A) over Blastoderma B. Fisch. & Breback 1809 (A) 1894 (A) and Rhodomyces Wettst. 1885 (A) The generic name Puccinia is typified by P. graminis, the and use rather than Aessosporon Van der Walt cause of the important wheat rust disease that has plagued 1970 (S) and Sporidiobolus Nyland 1950 (S)

84 IMA FUNGUS Asexual-sexual generic names in Pucciniomycotina and Ustilaginomycotina

The generic name Sporobolomyces, typified by S. be difficult to determine their affinities; the issue of genera ARTICLE salmonicolor, includes about 100 names and is widely typified by a sexual morphs is discussed further above (p. used. An earlier generic name, Rhodomyces, typified by 76). R. kochii, is now considered a synonym of S. salmonicolor The generic name Uromyces is already conserved over (Sampaio 2011). A second earlier monotypic generic name, Coeomurus Link ex Gray 1821 and Pucciniola L. Marchand Blastoderma, is typified by B. salmonicolor, the basionym 1829. of S. salmonicolor, thus these three generic names are synonyms. Another later name, Sporidiobolus, has six species, and is typified by S. johnsonii, a name now placed COMPETING SEXUAL AND ASEXUAL GE- in Sporobolomyces (Wang et al. 2015a). The generic name NERIC NAMES OF USTILAGINOMYCOTINA, Aessosporon, typified by A. salmonicolor, was described BASIDIOMYCOTA AND RECOMMENDATIONS as the sexual morph of Sporobolomyces salmonicolor (Van FOR USE der Walt 1970). Later authors showed that A. salmonicolor had the same heterothallic life-cycle and was conspecific Use Anthracoidea Bref. 1895 (S) rather than with S. salmonicolor (Fell & Statzell-Tallman 1980), thus Crotalia Liro 1938 (A) Aesseosporon and Sporobolomyces are synonyms. Among these five generic namesBlastoderma has rarely been used. The sexually typified genus Anthracoidea in Ustilaginales Although the name Rhodomyces appears in the literature, includes over 200 names, of which 90 are accepted in the three described species are not closely related to the Vánky (2012), while the asexually typified Crotalia has five type, as shown by Wang et al. (2015a). A second species of names, all of which also have names in Anthracoidea. The Aessosporon, A. dendrophilum, was described as the sexual type species of Anthracoidea is A. caricis. Initially the type morph of Bullera dendrophilum (van der Walt 1973), but is species of Crotalia, C. cintractiae-fischeri, was considered now regarded as Kwoniella dendrophila in Agaricomycotina to be the asexual morph of Cintractia fischeri; however, that (Liu et al. 2015), and unrelated to Sporobolomyces. Because species is now regarded as Anthracoidea fischeri (Vánky Sporobolomyces has priority, includes about 100 names and 2012). Although A. fischeri was not included in Hendrichs is widely used (GS Sporobolomyces = 10 600 Rhodomyces = et al. (2005), both A. caricis and A. fischeri are placed in 183, Blastoderma = 523, Sporidiobolus = 2970, Aessosporon = Anthracoidea by Vánky (2012), thus Anthracoidea and 171), we recommend that Sporobolomyces be protected over Crotalia are synonyms. Rhodomyces and Blastoderma, and that Sporobolomyces be Given its priority, the number of species, and more used rather than Sporidiobolus and Aessosporon. frequent citation in the literature (GS Anthracoidea = 619, Crotalia = 234), we recommend the use of Anthracoidea. Protect Uromyces (Link) Unger 1833 (S) over Uredo Pers. 1801 (A) Use Doassansia Cornu 1883 (S) rather than Savulescuella Cif. 1959 (A) The generic name Uromyces, typified by U. appendiculatus, includes over 800 names. When Persoon (1801) described The generic name Doassansia, typified by D. alismatis, Uredo, he included 30 species, one of which was U. betae. includes about 12 species (Vánky 2012) while Savulescuella, Laundon (1970) reviewed the history of the typification of the typified by S. alismacearum, includes only three names. generic name Uredo and concluded that U. betae should Vánky (2012) assumed that the asexual morph typified by be accepted as the type of the genus. Uredo betae is now S. alismacearum might be just “basidiospores of spores considered a synonym of Uromyces beticola (Boerema et al. that germinated in situ”, nevertheless Savulescuella was 1987, Simpson et al. 2006). Uromyces appendiculatus and validly published. Currently, S. alismacearum is considered U. beticola are regarded as congeneric, although species of a synonym of D. alismatis, making these names generic Uromyces are interspersed within Puccinia in Pucciniaceae synonyms. Given its priority, number of species, and (Aime 2006, Demers et al. 2017, Maier et al. 2003, Wingfield widespread use (GS Doassansia = 457, Savulescuella = 7), et al. 2004), thus Uromyces and Uredo are synonyms. Uredo we recommend the use of Doassansia. includes over 2000 names, but is not as widely cited as Uromyces (GS Uromyces = 26 400, Uredo 7500). Thus, we Use Doassansiopsis (Setch.) Dietel 1897 (S) recommend that Uromyces be protected over Uredo. rather than Doassansiella Zambett. 1970 (A) The generic name Uredo has been used for uredinial morphs of rust fungi belonging to multiple rust families, and The generic name Doassansiopsis is typified by D. does not reflect the affinities of species. Prior to 2011, this deformans. Sixteen additional names have been placed in generic name was used when a sexual morph was lacking and this genus, most of which are accepted by Vánky (2012). the species could not be described in the “correct” sexually The monotypic generic name Doassansiella is typified by D. typified genus according theCode even when it was possible aqualitis and has rarely been used. The type species is now to determine its generic placement. Now that placement of regarded as Doassansiopsis hydrophila (Vánky 2012). Given fungi in genera is no longer restricted by whether their types the priority, number of species, and more extensive use (GS are asxual or sexual morphs, in theory the need for the Doassansiopsis = 153, Doassansiella = 2), we recommend generic name Uredo no longer exists. However, in reality rust the use of Doassansiopsis. fungi are often found without the sexual morph and it can Doassansiopsis in Urocystales should not be confused

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with Doassansia and Doassinga in the Doassansiales be placed with confidence in any sexually typified lineage (Begerow et al. 2004). (Wang et al. 2015a; these species are therefore temporarily retained in Pseudozyma “pro tem.” Use Entyloma de Bary 1874 (S) rather than Entylomella Höhn. 1924 (A) Use Thecaphora Fingerh. 1836 (S) rather than ARTICLE Angiosorus Thirum. & M.J. O’Brien 1974 (S), The generic name Entylomella was first mentioned by Höhnel Rhombiella Liro 1939 (A) and Thecaphorella H. (1916), although in that publication a description is lacking of Scholz & I. Scholz 1988 (A) either the genus or species although one species, E. ranunculi, is listed. Later in discussing Cylindrosporium, Höhnel (1924) The type species of Thecaphora, T. hyalina, is now regarded mentioned Entylomella again as the conidial form of Entyloma as T. seminis-convolvuli (Vánky et al. 2008). The type and Doassansia species and two pages later provided a species of the asexually typified generic name Rhombiella, description of the type species as discussed by Braun (1995). R. cardamines, is now regarded as Thecaphora thlaspeos Entylomella ranunculi is the type of Entylomella, which is (Vánky et al. 2008). The type of Thecaphorella, T. antherarum, now regarded as a synonym of Entyloma ficariae. The type was described as the asexual morph of Thecaphora seminis- species of Entyloma is E. microsporum. Begerow et al. (2002) convolvuli (Scholz & Scholz 1988). Begerow et al. (1997) demonstrated that E. ficariae and E. microsporum belong in demonstrated that Thecaphora represents a well-resolved Entyloma, thus Entylomella and Entyloma are synonyms. genus in Ustilaginales including the types of T. seminis- Over 400 names are listed in Entyloma and this generic convolvuli and T. thlaspeos; these three generic names are name is well known (GS = 2410) while 59 names have been therefore synonyms. Similarly, Andrade et al. (2004) and described in Entylomella, many of which have been placed Vánky et al. (2008) demonstrated that Thecaphora includes T. elsewhere, and the name has been used infrequently (GS seminis-convolvuli and T. solani (syn. Angiosorus solani, type = 60). Given the priority, greater number of names, and of the monotypic generic name Angiosorus), an economically familiarity, we recommend the use of Entyloma. important fungus causing potato smut (Mordue 1988). Vánky (2012) includes over 60 species in Thecaphora, under which Use Mycosarcoma Bref. 1912 or Ustilago (Pers.) he lists four sexually typified generic names as later synonyms, along with the rejected name Sorosporium. Thecaphora has Roussel 1806 (S) rather than Pseudozyma priority, includes numerous species, and is well known (GS Bandoni 1985 (A) Thecaphora = 1420, Rhombiella = 5, Angiosorus = 128), thus The yeast-like morphs of Ustilaginales have been described we recommend the use of Thecaphora. in the genus Pseudozyma. Boekhout (1995) showed that the type species of Pseudozyma, P. prolifica, was closely Use Tilletia Tul. & C. Tul. 1847 (S) rather than related or synonymous wth Ustilago maydis, the cause of Tilletiella Zambett. 1970 (A) corn smut (Ainsworth 1965). While 26 names have been described in Pseudozyma, many of these have now been The genus Tilletia has long been accepted as the name for placed in diverse sexual genera such as Anthracocystis, bunt fungi with over 100 accepted species, some of which Dirkmeia, Kalmanozyma, Langdonia, Moesziomyces and cause serious diseases of crops in the grass family such as Sporisorium. A controversy exists over which generic name common bunt and dwarf bunt of wheat (Hoffman 1982, Mathre to use for the species long referred to as U. maydis, an 1996). Tilletia, typified by T. caries, and treated as T. tritici, is earlier name for P. prolifica. The generic name Ustilago now placed in Tilletiaceae (Tilletiales, Exobasidiomycetes; had been considered to be typified by U. hordei, cause of Begerow et al. 2006). The monotypic Tilletiella was established barley smut (Clinton 1906, Vánky 2012). This and related as an asexually typified genus forT. alopecuri, but that species species, such as U. avenae, U. striiformis, and U. tritici is now regarded as Tilletia alopecuri (Vánky 2012). Tilletiella is were determined by McTaggart et al. (2016) to fall in a clade therefore a synonym of Tilletia. Given the priority, large number distinct from the one that included U. maydis, type of the of species, and widespread use (GS Tilletia = 16 900, Tilletiella resurrected generic name Mycosarcoma, which was based = 0), we recommend the use of Tilletia. on morphological and cultural characteristics by Brefeld (1912). Based on molecular data, five names were included Use Urocystis Rabenh. ex Fuckel 1870, nom. in Mycosarcoma by McTaggart et al. (2016), resurrecting cons. (S) rather than Paepalopsis J.G. Kühn Mycosarcoma maydis for the corn smut and closely related 1882 (A) species. However, a proposal has been made to conserve the name Ustilago with U. maydis as the conserved type The type species of Paepalopsis, P. irmischiae, is considered (Thines 2016). Pseudozyma prolifica is a name used for the the asexual morph of Urocystis primulae, a species included in asexual morph of Mycosarcoma / Ustilago maydis, thus we Urocystis, which is typified byU. occulta (Vánky 2012); these recommend the use of Mycosarcoma or Ustilago depending generic names are consequently synonyms. Vánky (2012) on whether the NCF votes to conserve the name Ustilago listed four sexually typified generic synonyms of Urocystis, with a new type. Because the generic name Pseudozyma including Tubercinia Fr. 1832 nom. sanct. and Polycystis is no longer available, the taxonomic treatment of several Lév., earlier names of Urocystis which has been conserved names in Pseudozyma (e.g. P. alboarmeniaca, P. thailandica, over them (Wakefield 1939). Three names have been placed P. tsukubaensis, P. pruni) is unclear as they cannot currently in Paepalopsis, two of which are now regarded as species

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of Urocystis, while the third, P. deformans, is now placed in Begerow D, Stoll M, Bauer R (2006) A phylogenetic hypothesis ARTICLE the ascomycete genus Hapalosphaeria as H. deformans. of Ustilaginomycotina based on multiple gene analyses and Vánky (2012) accepted over 160 species of Urocystis in his morphological data. Mycologia 98: 906–916. world monograph of smut fungi. Given the priority, number Berndt R (2015) The rust fungi (Uredinales) on ferns in South Africa. of species, and widespread use (GS Urocystis = 4920, Mycological Progress 7: 7–19. Paepalopsis = 24), Urocystis remains the correct name for Boekhout T (1995) Pseudozyma Bandoni emend Boekhout, a genus this genus. for yeast-like anamorphs of Ustilaginales. Journal for General Applied Microbiology 41: 359–366. Boerema GH, Loerakker WM, Hamers MEC (1987) Checklist for ACKNOWLEDGEMENTS scientific names of common parasitic fungi. 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ARTICLE Wright JE, Bandoni R, Oberwinkler F (1981) Agaricostilbum: an auricularioid basidiomycete. Mycologia 73: 880–886. Yun HY, Hong SG, Rossman AY, Lee SK, Lee KJ, et al. (2009) The rust fungus Gymnosporangium in Korea including two new species, G. monticola and G. unicorne. Mycologia 101: 790–809.

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